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-8 9 -
HEPAT1CS WITH HAND-LENS.
A. J. Grout.
There has been a considerable demand for a simple book on the
Hepatics. To meet this demand I am preparing a treatment of the Hepaties
similar to that which I have given the mosses in "Mosses with a Hand-
Lens." This will be included in the second edition of that book now in pre-
paration (See adv. in this No. of the Bryologist). This key to the genera is
printed here with the hope that it will be used and criticised by the readers
of the Bryologist, and that by the assistance of these criticisms the final
treatment may be made more helpful. With the Queen ^-inch achromatic
triplet I am able to make out the more minute structures mentioned in the
keys. Many of them, especially leaf structure, can not be made out satisfac-
torily unless the objects be mounted in water on a slide in the same man-
ner as for a compound microscope. The slide should then be held up to the
strong light, the slide being held with the left hand and the lens with the
right, the right thumb resting upon that of the left hand so that the focus
will not be distributed by any unsteadiness of the hands.
From now until winter closes in I shall be glad to attempt to name
Hepatics for our subscribers if the speciments be accompanied by a stamp,
full data for the label, and the best name the collector can give. Fresh
material only is desired. Almost none of the books give the time of matur-
ing spores of the different species, and I hope that our readers will send me
all the data of this sort that they have. Comparatively few illustrations are
possible in this article, but the figures in the sixth edition of Gray's Manual
will prove very helpful. In working up this key I have been surprised to
find that sterile Hepatics are, as a rule, much easier to identify than sterile
mosses. Many of the species maturing their spores in early spring have
the spores and capsules pretty fully developed in the preceding autumn so
that some of the sporophyte characters are nearly- always accessible.
Hepatics shrivel more than mosses in drying and are best studied while fresh,
especially the thalloid forms.
A few of the rare genera are omitted and in the completed treatment
some of the minute or difficult species will not be included.
The Germans call the true mosses Laubmoose, meaning leafy mosses,
and the Hepatics, Lebermoose, or liver mosses, The name Liverwort was
originally applied to Marchantia because of its fancied resemblance to the
liver. Because of this resemblance it was supposed to be a specific for all
liver troubles according to the old doctrine of signatures. From this came
the Latin name Hepaticae and the German Lebermoose. "Thus does the
language of ignorant superstition become the adopted language of science."
The chief distinctions between Mosses and Hepatics have been noted
in the Bryologist for April, 1899, but a few additional notes here may prove
The Hepatics may be leafy stemmed and appear much like mosses, or
they may consist of a broad, flat and rather thin stem (thallus) which is
usually closely applied to the substratum. These thalloid Hepatics might
be mistaken for some of the foliaceous lichens but the Hepatics are always
much greener and produce spores in a very different manner.
In the leafy-stemmed Hepatics, often called Scale Mosses, the leaves are
without midrib and are nearly always in two ranks and flattened so as to lie
in one plane, but in the great majority of cases there is a third rudimentary
row on the inner side which are called underleaves, or amphigastra by those
devoted to technical names. The pedicel which corresponds to the seta of
the mosses does not, as a rule, grow much until the spores are nearly ripe,
when it elongates very rapidly. The pedicels and capsules are of a much
more delicate structure than in the mosses so that they disappear soon after
the spores have escaped, but the peculiar and characteristic scales or bracts
around the base of the pedical often remain much longer and help greatly in
identifying species. Immediately surrounding the base of the pedicel is a
tubular, somewhat three-sided organ called the inner involucre or perianth,
surrounding this the outer involucre, called simply involucre by many
authors. This latter may be either tubular or composed of separate leaf-like
divisions of varied shapes, called involucral leaves or bracts, or perichatial
leaves or bracts, or simply bracts. Either one, or even both, of these invo-
lucres may be lacking in some species.
So far as possible gametophyte characters have been used in the keys
and descriptions and in the great majority of cases identification is easy from
this part of the plant alone. Hepatics generally grow in moist situations on
soil, roots of trees, and decaying wood.
Key to Families.
Plants leafy, mosslike in appearence except for the two-ranked leaves
entirely lacking midrib Scale Mosses (Jungermanniaceee) .
Plants consisting of a flattened green thallus, sometimes nearly circular but
usually elongated and branching. (See illustrations of Riccia, Mar-
chantia, Anthoceros, etc.) A.
i. Capsules, if present, immersed in the tissue of the plant. Plants float-
ing on the surface of still water or floating on the mud along the
Capsules raised well above the thallus. Plants often growing in mud but
never floating 2 -
2. Stomata (in our genera) present, easily discernable with a lens as small
pores on the upper surface of the rather thick thallus ; capsule borne on
a special stalked receptacle as in Marchantia.
Stomata not present, on the thinner thallus: capsules never borne on a
special stalked receptacle 3-
3. Capsules very long and slender, splitting into two valves when ripe after
the manner of a mustard pod, the slender hairlike columella remaining
in the center Horned Liverworts (Anthocerotaceae).
Capsules globular or ovoid, splitting into four valves; columella lacking.
Thalloid Scale Mosses (Metzgeriaceag).
Fig. i. a, Sterile and b, fertile thallus Anthoceros punctatus X 2 & 1.
Fig. 2-6. Marchantia polymorpha, from Bryolouist, 4:34-35, 1901. Fig. 2.
Male plant a little reduced, showing antheridial receptacles. Fig. 3.
Longitudinal section of antheridial receptacle magnified. Fig. 4.
Female plant reduced showing the stalked receptacles which characterize
this family. These receptacles vary in the family from the shape shown
in this figure to almost perfectly conical and entire. Fig. 5. Section of
a part of a female receptacle magnified, showing two sporogonia. The
seta of one has elongated, pushing the capsule out from the outer fringe
(involucre) and the inner fringe (perianth) at the base of the seta is a lit"
tie collar representing the base of the broken calyptra. Fig. 6. Sterile
thallus with gemma;.
THE TRUE LIVERWORTS (MARCHANTIACE/E).
The plants of this family consist of thallus of medium to large size, one-
half to six inches in length, usually branching dicotomously but sometimes
with more than two branches at a fork. They are attached to the substratum
by numerous roothairs and are thickened in the middle to form a midrib.
This in some cases is not vary apparent above but shows plainly underneath.
The upper surface is covered with small pores (stomata) which are very
apparent with a lens, except in Reboulia. The capsules are spherical or ovoid
and open irregularly by imperfect valves or by a portion of the top coming
off after the manner of a lid. In this family the capsules and usually the
antheridia are borne on special long-stalked receptacles well illustrated by
the familiar Marchantia.
Key to the Genera.
i. Sterile stems bearing abundant gemmae in shallow open receptacles. . . .2
Sterile stems without gemmae 3
2 Found only in and around greenhouses: gemmae in crescent-shaped
receptacles ; never fruiting in our region Lunularia.
Growing abundantly everywhere; gemmae in cup-shaped receptacles: cap-
sule-bearing receptacle with 7-11 conspicuous rays Marchantia.
3. Thallus large ; 2-6 inches long and l / 2 inch or more wide, distinctly areo-
late as in Marchantia, but areolas larger and hexagonal. . . Conocephalus.
Thallus less than two inches in length and much narrower 4
4. Pores (stomata) scarcely distinguishable ; antheridia in sessile receptacles
which might be mistaken for gemmae; thallus purple on the margins;
midrib strong underneath but not conspicuous above Reboulia.
Pores conspicuous, white; antheridia in peduncled disk-like receptacle; thal-
lus with numerous dark purple scales underneath Preissia.
Pores conspicuous; antheridia immersed in the thallus; thallus purple
underneath, at least along the margins 5
5. Perianth conspicuous, split into 8-16 fringe-like lobes; peduncle not
Perianth lacking: peduncle chaffy at top and bottom Grimaldia.
(The Reboulia of this key is the Asterella of Gray's Manual and the
Asterella is the Fimbriaria of that work. )
THE THALLOID SCALE MOSSES (METZGERIACE/E).
The spore bearing portion of plants of this family is like that of the
Scale Mosses, but the green part of the plant is a thallus instead of a leafy
stem in nearly all cases. There are, however, some intermediate forms in
the family in which the thallus is divided into leaflike lobes. The thallus is
much less highly differentiated than in the Liverworts and Riccias: there
are no areolae or pores (stomata), and the thallus is much thinner than in the
Liverworts, in some species consisting of only a single layer of cells except
at the midrib. The capsules are borne singly on setae arising directly from
the thallus. They are spherical to elongated-ovoid and remain enclosed in
the calyptra until mature when the seta? rapidly elongate and break open the
calyptra which is left at the base of the seta. The capsules open by four
valves as in many of the Scale Mosses. A careful search of wet bare earth
in shaded or springy places will nearly always yield one or more species of
Key to the Genera.
1. Thallus with a distinct midrib 2.
Thallus without a distinct midrib 4-
2. Thallus 1/25 to 1/12 inch wide, dichotomously branched, cilliate along
the margins Metzgeria.
Thallus ^ to y 2 inch wide, not cilliate at margins, entire or lobed 3.
3. Thallus simple or only once forked, 1 to 4 inches long, prostrate: margins
sinuate to entire: capsule ovoid-cylindric Pallavicinia.
Thallus dichotomously branched, j^ to 1^ inches long, often densely clus-
tered and ascending, margins lobed: capsules spherical without perianth,
^^ appearing buried in the midrib for
•v. ^i ®wJf£\ sft> some time before the ripening of spores
^Mtf! W^^^^ML* (Fig- 7) Blasia.
vw 1 ^a^k6^^^^^ ■*• Thallus pinnately or palmately branch-
\w?-- n^i/ft; e< ^' J / 24 to I//I2 * n w 'de (except
)ffy a Fi'WJfr b R. pinguis) Riccardia.
•^Sg Thallus subsimple or dichotomously
branched, y& to 'A, inch in width (Fig.
tlg - 7 - 8) Pellia.
Fig. 7. Blasia pusilla L. a. Fertile plant in August
showing capsule in position. At the side is shown the cap-
sule removed from the thallus. b. Sterile plant with flask-
shaped bodies which produce gemmae.
,,,, v . Fig. 8. Pellia epiphylla Raddi. Thallus X 1, showing
involucre and position of capsule as it appears in August.
THE SCALE MOSSES (JUNGERMANN1ACE/B).
The reproductive part of the Scale Mosses, including the ripened cap-
sule and its connected parts, perianth, involucre, etc., is essentially as in
the Thalloid Scale Mosses, but the vegetative part strongly resembles the
true mosses in general appearance. The leaves, however, are apparently
flattened out into two rows, one on either side of the stem. They are
entirely without midrib and are frequently two-cleft or lobed. One of the
lobes is often smaller and folded under the other making the leaves " com-
plicate-bilobed," in the language of the books as shown in the illustrations
of Radula and Porella. This can best be made out by holding a single
stem up to the light and examining with a lens, when the under lobe will
show plainly as a deep shadow. In Scapania, the under lobe is the larger
and the plants look as if there were four rows of leaves. The lower lobe is
called the lobule and the upper simply the lobe. Very many species have a
third rows of leaves on the under side of the stem called technically " amphi-
gastra" or underleaves, these vary in size from one-third the size of the
ordinary leaves to so minute that high powers of the compound microscope
are needed to see them clearly. The upper margin of the leaves may over-
lap the lower margins of the leaves next above as in Porella, or the upper
margin of a leaf may lie under the lower margin of the leaf next above as
in Plagiochila. In the former case the leaves are said to be incubous, in
the latter succubous. As this distinction is in most cases easy to observe,
it is given a prominent part in the key. Occasionally the leaves are so far
apart that it is hard to determine the leaf arrangement, but a careful search
will usually discover some plants in which this character can be seen.
In plants with incubous leaves the bud is turned downward: when the
leaves are succubous the terminal bud is turned up. So far as possible the
key has been based upon the leafy or vegetative portions of the plants, but
in some few cases the characters connected with the reproductive organs
and capsules are necessary to accurately determine a plant. In most cases
the characters used can be determined without mounting, if, however, they
can not be readily made out the parts should be mounted as for the com-
pound microscope. If one has access to a compound microscope itwill often
prove a very great help, although not necessary to make out the characters
mentioned. Mnium and Fissidens are sure to be mistaken for Hepatics by
the beginner unless the midrib or the leaves is noted.
Key to the Genera and Species.
i. Leaves entirely or in large part composed of hair-like divisions (easily
observed if held up towards a strong light) 2.
Leaves not as above 3.
2. Plants grayish green, growing over the ground amid mosses in cool
bogs, at least twice pinnate and somewhat resembling the Fern Mosses;
leaves divided to base into hair-like lobes Trichocolea.
Plants dark green, much smaller, growing chiefly on rotton wood, but also
found on humus-covered stones and soil ; leaves with a considerable
solid portion Ptilidium.
Plants exceedingly minute, looking like a small green alga or moss proton-
ema. Scarcely recognizable except when fruited; common on decayed
wood, moist soil, etc Blepharostoma.
3. Leaves incubous A.
(Scapania and Chiloscyphus forms may be sought here.)
Leaves succubous B.
1. Leaves complicate-bilobed, upper lobes entire or nearly so (except
Jubula). See figures and description of Porella 2.
Leaves sometimes lobed or cleft but not complicate-bilobed 5.
2. Plants blackish or brownish green, minute, leafy stems 1/25 inch or less
wide; lobule like an inflated sac (Plate XIV) Frullania.
Plants often dark olive-green but not often blackish; 1/16 inch in width,
lobule not sac-like 3.
3. Under leaves lacking; perianth strongly flattened crosswise (Plate
Underleaves conspicuous 4 .
4. Lobule with its longer edge attached to lower margin of lobe (See cuts,
Plate XIII) Lejeunea.
Lobule with its shorter margin attached to the lower edge of lobe (Plate
5. Leaves mostly entire Kantia.
Leaves strongly toothed, notched, or cleft at apex 6.
6. Leafy stems less than 1/25 inch in width Lepidozia.
Leafy stems 1/16 to % inch in width, with downward growing stolons
(Plate XIII) Bazzania.
1. Leaves complicate-bilobed, lobes nearly equal or the lower larger giv-
ing the appearance of four rows of leaves of which the two upper are
incubous and the two lower succubous Scapania.
Leaves not complicate-bilobed, in some cases toothed or divided 2.
2. Leaves undulate on the margin; plants densely clustered; roothairs
bright claret colored Nardia hyalina.
Leaves entire or slighly emarginate ; roothairs colorless 3-
Some or usually all of the leaves strongly toothed or lobed; roothairs color-
3. Leafy stems at least ]/% inch wide, leaves plainly overlapping, on ground
and over mosses 5-
Plants about y% inch wide: many leaves not overlapping 4-
Leafy stems 1/16 inch wide or less 6.
4. Aquatic, floating, underleaves absent.
Chiloscyphus polyanthus var. rivularis.
On old logs and moist ground, underleaves present Chiloscyphus.
5. Plants creeping; leaves oblong to oblong-ovate, decurrent — Liochlaena.
Plants ascending; leaves round obovate, not decurrent Plagiochila.
6. Leaves with a border of larger cells which appear as a whitish margin
under the lens Odontoschisma and Nardia crenulata.
Leaves without border of larger cells Jungermannia Schraderi.
7. Upper leaves with a strongly many toothed margin Plagiochila.
Leaves 3-5 cleft Jungermannia barbata.
Leaves two toothed or cleft 8.
8. Plants minute, leafy stems less than 1/25 inch wide ; underleaves absent
or so small as to be invisible with a lens: leaves round-ovate to
obovate, cleft for at least ]^ their length Cephalozia.
(Some small species of Jungermannia may be sought here but their
leaves are less deeply cleft and the plants are a much darker green).
Leafy stems at least 1/16 inch wide: leaves two toothed but scarcely cleft .9
9. Underleaves % the size of the other leaves Lophocolea minor.
Underleaves absent or minute 10.
10. Leaves subvertical, varying from bidentate to refuse, or even entire
near apex Lophocolea heterophylla.
Leaves with the edge attached nearly lengthwise of the stem, extending out at
almost right angles from it and lying flat in a horizontal plane. Geocalyx.
Leaves subvertical and all alike inserted more nearly crosswise of the
stem 11 -
n. On sterile ground in open woods Jungermannia excisa.
On rotton wood I2 -
12. Leaves round-ovate with an obtuse sinus between the teeth. Harpanthus.
Leaves subrectangular with an acute sinus; plants dark to brownish
g re en Jungermannia Michauxii.
Plate XIII. Bazzania, Radula, Lejeunea.
Plate XIV. Porella, Frullania.
- 9 8-
EXPLANAT10N OF PLATES.
Plate XIII. Fig. 9. Bazzania trilobata (L.) S. F. Gray. From Bryologist,
4:68, 1901. A. Plant slightly magnified showing flagella springing
from the underside. B. (1.) Portion of female plant with capsule. (2.)
Capsule open. C. Involucre, perianth and base of seta enlarged.
The involucre consists of the small leaves at the bottom of the fig-
ure. D. Male plant seen from below, showing antheridial branch,
minute underleaves and incubous arrangement of leaves. E. & F.
Illustrate spiral elators, spores and cell structure of leaf which cannot
be seen clearly with a hand-lens.
Fig. 10. Radula complanata Dumort. A. Plant natural size. B. Branch
with fruit showing clearly the seta and capsule, with the calyptra at
base of seta showing through the transparent tubular perianth, and at
base of the perianth, the involucre. This misrepresents the leaves,
making them appear succubous. C. Leaf showing lobule with roothairs
and larger lobe with gemmae along the edge. This illustrates the sim-
plest forms of " complicate-bilobed " leaf. There are no underleaves.
D. Calyptra. E. Spores, highly magnified.
Fig. 11. Various species of Lejeunea, from Bryologist, 6:-27, 1903.
Showing underleaves in all but the right hand figure. Note that the
lobule is attached to the lobe by its longer edge.
Plate XIV. Fig. 12. Porella pinnata L. From the Bryologist, 5: 34,
1902. A. Underside of stem showing narrow underleaves and narrow
lobules attached by their shorter edge to lobe. B. Single leaf showing
lobe and lobule.
Fig. 13. Porella platyphylla (L.) Lindb. From Bryologist, 5: 35, 1902.
B. Upper side of stem showing perianth and emerging capsule. Also
showing clearly incubous arrangement of leaves. C. Underside of
stem, the leaves shown too far apart. D. Longitudinal section of
perianth. E. Capsule. F. Leaf. G. Part of plant showing male
Fig. 14. Frullania. From Bryologist, 5 : 4, 1902. I. Plant of Frullania
Eboracensis Gottsche., on the bark of birch. II. Underside of same
showing underleaves and the queer saclike inflated lobules which
remind one of the bladders of Utricularia. III. and IV. Under and
upper side of F, Asagrayana Mont. V. Involucre and perianth of F.
Fig. 15. Ptilidium ciliare (L.) Nees. a. Leaf X 37. b. Plant with
perianth and young capsule X 2. c. Portion of plant X 5.